Plants employ diverse defense mechanisms to combat attack by harmful organisms. For instance, plants produce constitutive physical barriers or use chemical compounds such as specialized secondary metabolites to resist herbivore or pathogen invasion. Considering the cost-efficiency and energy balance between defense, growth and reproduction, defense reactions in plants have to be regulated temporally and spatially. As more cost-efficient strategies, plants may induce their defense response only in the presence of the attacker or restrict constitutive defenses to specific tissues or cells. In this study, we investigated aspects of the spatial regulation and induced changes of primary and secondary metabolism in roots of Arabidopsis thaliana. Roots represent important organs for anchoring plants in the soil and taking up water and nutrients. Hence, it is assumed that roots are as well protected as aerial tissues by different defense mechanisms. The first part of this work is focused on the cell-type-specific biosynthesis of volatile terpenes in Arabidopsis roots. Terpenes are the most abundant specialized metabolites in plants and play an important role in plant defense against pathogens or herbivores. Terpene biosynthetic enzyme activities are often coordinated in specific tissues and cellular compartments. Fine-scale transcriptome maps of Arabidopsis roots have shown that terpene biosynthesis is restricted to particular cell types. However, the reasons and significance of this cell-type specificity are not well understood. We hypothesized that the formation of terpene metabolites is not restricted to specific cells but can be supported by different cell types. We, therefore, probed the plasticity of the cell-specific formation of terpenes by swapping the expression of the terpene synthase (TPS) genes, TPS08, TPS13 and TPS25, between different root cell types in the respective mutant background. To investigate the ectopic expression of TPSs at different levels, quantitative real-time PCR (qRT-PCR), confocal microscopy, and gas chromatography-mass spectrometry (GC-MS) were performed. We found that terpene synthase TPS08, which produces the diterpene rhizathalene and is normally expressed in the root vascular tissue, is functionally active when expressed in the epidermis or cortex, although at substantially lower levels compared to the wild type. We did not find an obvious correlation between the volatile emission level and gene transcript level of TPS08, which may be attributed to a reduced activity of the expressed TPS08-yellow fluorescent protein (YFP) fusion protein. When expression of TPS13 (producing the sesquiterpene (Z)-"-bisabolene) was directed from the cortex to the epidermis or stele, TPS13 gene expression and (Z)-"-bisabolene formation was supported by these cell types although to varying levels in comparison to wild type. TPS13-YFP fluorescent signal driven by the epidermal WER and GL3 promoters was primarily detected at the root tip. Terpene production was also observed for the (E)-"-farnesene sesquiterpene synthase TPS25 when its expression was switched from the endodermis and non-hair producing epidermal cells to hair producing epidermal cells although only a weak fluorescent signal was detected from the expressed TPS25-mGFP protein. Together, the results provide preliminary evidence for a relaxed cell specificity of terpene biosynthesis in Arabidopsis roots and suggest that tissue-specific terpene metabolite patterns could change depending on different selective pressures in rhizosphere. In the second part of this study, we performed global gene transcript profiling and primary metabolite analysis of Arabidopsis roots upon feeding by the generalist root herbivore, Bradysia (fungus gnat). In a microarray analysis, we identified 451 of 22,810 genes that were up-regulated more than 2-fold. Gene ontology (GO) analysis showed that 26% of those genes with predicted or known functions play a role in primary or secondary metabolism, while 24% are involved in cell signaling or in responses to stimulating factors, such as jasmonic acid (JA), ethylene, wounding, and oxidative stress. At the metabolite level, we observed only marginal changes of amino acid, sugar and carboxylic acid relative levels over a time course of 4 days of Bradysia feeding. There was a trend for increased levels of amino acids and the relative levels of sucrose were increased significantly ("=0.05) at the fourth day of feeding. In conclusion, the study provided evidence for the induction of genes related to primary and secondary metabolism and stress responses in Arabidopsis roots, but showed only marginal changes at the primary metabolite level. In addition, the work indicated that the formation of terpene-specialized metabolites in Arabidopsis roots is not restricted to specific cells, but can be supported by different cell types. / Master of Science

AN ABSTRACT OF THE THESIS OFNicholas Seaton, for the Master of Science degree in Plant Biology, presented on December 4, 2019 at Southern Illinois University Carbondale. TITLE: THE EFFECT OF DEER BROWSE ON ACHYRANTHES JAPONICAMAJOR PROFESSOR: Dr. David GibsonPlants respond in many ways to damage. These responses vary between sites depending on the severity and duration of the incident. One common form of damage in the forest understory is herbivory or browse. White tailed deer (Odocoileus virginianus) have been observed to change the dominant species of forests by selectively browsing palatable species in the understory. These changes in species dominance can lead to unwanted consequences, sometimes resulting in a proliferation of weedy or invasive plants or a reduction in performance and competitive abilities based on morphological traits. Understanding the changes that occur to undesirable species after deer browse can help land managers in their prioritization of sites for land management and understand the driving forces behind a species’ success or failure. Using deer exclosure plots, this study looks at the effects of white-tailed deer on Achyranthes japonica, an herbaceous invasive species in the Ohio River floodplain of Illinois and surrounding states. White tailed deer have been observed to browse A. japonica throughout the invaders range, but little is known about the plant’s response. Deer browse data were collected in the summer of 2018 from May to August. Estimated deer densities among six study sites ranged from 8 to 22 deer per km2. Plants that were browsed during the growing season were morphologically different to those that were not browsed. Browsed plants were 11.5 ± 0.1 cm shorter (F1,218=11.658; p<0.001) on average and produced 0.33 ± 0.09 fewer nodes (F1,216= 4.045; p<0.05). Browsed plants also produced 2.7 ± 0.32 fewer flowering spikes and were similar in length to those of un-browsed plants. These morphological differences showed significant variation between sites. Floristic Quality Indices of the herbaceous plant communities (Ȳ =3.5) ranged from 3.2 to 3.9 among study sites. This study shows that site conditions can impact the response of A. japonica growth as it continues to invade across its current introduced range and that the species is adaptive and grows along-side other similar weedy species such as Microstegium vimineum and Parthenocissus quinquefolia. This study also indicated that deer browse has little to no impact on the growth of Achyranthes japonica.Keywords: Achyranthes japonica, Odocoileus virginianus, herbivory, browse, deer density, site quality

In Europe, fossil fruits and seeds of Rhodoleia (Hamamelidaceae) have been described from the Upper Cretaceous to the Miocene, whereas no fossil record of Rhodoleia has been reported in Asia, where the modern species occur. Herein, 21 fossil leaves identified as Rhodoleia tengchongensis sp. nov. are described from the Upper Pliocene of Tengchong County, Yunnan Province, Southwest China. The fossils exhibit elliptic lamina with entire margins, simple brochidodromous major secondary veins, mixed percurrent intercostal tertiary veins, and looped exterior tertiaries. The leaf cuticle is characterized by pentagonal or hexagonal cells, stellate multicellular trichomes, and paracytic stomata. The combination of leaf architecture and cuticular characteristics suggests that the fossil leaves should be classified into the genus Rhodoleia. The fossil distributions indicate that the genus Rhodoleia might originate from Central Europe, and that migrated to Asia prior to the Late Pliocene. Additionally, insect damage is investigated, and different types of damage, such as hole feeding, margin feeding, surface feeding, and galling, are observed on the thirteen fossil leaves. Based on the damage frequencies for the fossil and extant leaves, the specific feeding behavior of insects on Rhodoleia trees appears to have been established as early as the Late Pliocene. The high occurrence of Rhodoleia insect herbivory may attract the insect-foraging birds, thereby increasing the probability of pollination.

Acropora cervicornis, commonly known as the staghorn coral has historically been a major contributor to reef structural complexity, providing habitat for many functionally important fish species throughout Florida and the Caribbean. Unfortunately, due to disease, bleaching, and local anthropogenic stressors, A. cervicornis populations have suffered drastic declines that have negatively impacted associated reef fish populations. In order to promote recovery, A. cervicornis fragments can be cultivated in nurseries and outplanted back onto reefs. This practice can effectively increase A. cervicornis abundance, but the long-term effects on local fish assemblages, and specifically functionally important grazing fishes, has not been assessed. Fish assemblages at natural (control) sites were compared to outplanted A. cervicornis sites in Southeast Florida. Fish surveys were conducted each summer at four locations from 2012 to 2017. Each location contained three outplanted A. cervicornis and one or two control sites. Outplant sites were defined by 50 A. cervicornis colonies in a 36 m2 area. Control sites occupied the same area but did not contain outplanted colonies. The fish assemblage structure was assessed in terms of composition, demography, and functional temporal trends as well as with the increasing structural complexity of the outplanted corals, defined as total linear extension (TLE). Significant temporal trends were recorded for total fish abundance, grazer abundance, and diversity. Structural complexity (outplanted A. cervicornis measured in TLE m-2) was found to be a significant predictor of total fish abundance, grazer abundance and diversity. Fishes 2-5 cm total length were most numerous indicating that the outplant sites may be providing habitat for juvenile reef fishes, particularly algae consumers. These findings suggest that A. cervicornis restoration may be creating a positive feedback loop in which outplanted corals create habitat for grazing fishes that in turn reduce algae competition, potentially providing new habitat for coral settlement.

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Coastal wetland plants serve as ecological engineers in a physiologically stressful environment and the loss of coastal wetlands can cause negative effects throughout the estuarine system. Due to increased degradation of coastal habitats worldwide, interest in restoration has increased around the world. An understanding of the biotic processes affecting species distribution and diversity is critical for future conservation, management, and restoration of coastal wetlands. The purpose of my study was to test the effects of biotic interactions on native coastal wetland plants and determine how these interactions may be incorporated into current and future restoration projects. I had three primary goals for my dissertation. First, I evaluated the effectiveness of natural regeneration of coastal wetland communities following hydrological restoration. Second, I examined effects of biotic interactions between mangroves and other wetland species by experimentally testing: 1) trapping capabilities of early successional plant species on Rhizophora mangle propagules, 2) effects of pre-dispersal propagule damage on native mangrove species, 3) facilitative and competitive interactions between the plants Rhizophora mangle, Batis maritima, Sarcocornia perennis, and the fiddler crab, Uca pugilator using mesocosms. Third, I evaluated the combined effects of biotic and abiotic interactions on survival and growth of R. mangle propagules during their first year of establishment using a manipulative field experiment. Results from my study increase our knowledge of the importance of biotic interactions in coastal wetland communities, their role in early successional stages, and have direct applications to coastal wetland restoration and management.

Mammalian herbivory of bottomland hardwood seedlings has been listed as one of the primary causal factors of failed afforestation efforts in the Lower Mississippi Alluvial Valley (LMAV). This study examined the biological and economic effects of mammalian herbivory on recently afforested carbon sequestration sites in the LMAV. Selected seedlings of six planting mixes were observed through the first year following planting to monitor seedling survival, growth, and mammalian herbivory. It was determined that greater than 10% of selected seedlings were browsed by various mammalian herbivore species, and some species mixes were browsed in excess of 50%. Financial analyses compared alternative afforestation strategies and determined to what extent herbivore-induced seedling mortality could reduce investment returns of landowners engaged in afforestation activities. In the presence of extreme mammalian herbivory, landowner returns can be reduced by hundreds of dollars per acre and could prevent further afforestation activities in the LMAV.

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